Microsyops | |
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Order: | Plesiadapiformes |
Family: | † Microsyopidae |
Subfamily: | † Microsyopinae |
Genus: | † Microsyops Leidy, 1872 |
Species | |
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Microsyops is a plesiadapiform primate found in Middle Eocene in North America. [1] It is in the family Microsyopidae, a plesiadapiform family characterized by distinctive lanceolate lower first incisors. [2] It appears to have had a more developed sense of smell than other early primates. [3] It is believed to have eaten fruit, and its fossils show the oldest known dental cavities in a mammal. [4] [5]
There are nine species of Microsyops that exist in the fossil record from the middle Wasatchian (~53 million years ago) through Uintan (~42 million years ago) North American Land Mammal Ages. Microsyops is primarily known from the Rocky Mountain region of the United States, though fossils have also been found in California and Texas. [6] [7]
Body size diversity of Microsyops spans from the 700-gram Microsyops cardiorestes to over 3000 grams for Microsyops kratos, estimated using the dimensions of the upper and lower last premolar and first molar. [8] [9] The diet of Microsyops is varied among the nine species, with the smaller-bodied and more primitive species, like M. cardiorestes, likely able to exist by eating almost exclusively insects. [9] However, larger-bodied species, such as Microsyops annectens or M. kratos, likely needed to expand their diets to include other food sources. This is also supported by wear facets on the molars of M. annectens and M. kratos that are indicative of heavier shearing and crushing forces required of harder foods like fruits and nuts. [9] Another indication of expanded diets, away from strict insectivory, comes in the form of reported cavities in a sample of Microsyops latidens where a sample of 1030 individuals included 77 specimens showing signs of cavities. In this case, cavities are likely caused by a reliance on more sugary foods, such as fruits, moving away from strict insectivory. [10]
Consistent with other North American members of Microsyopidae, Microsyops has a lower central incisor that is enlarged, procumbent, and lanceolate. The expansive flattened surface of the lower central incisor is oriented towards the front of the tooth. [2] Microsyops has a lower dental formula of 1-0-3-3, with one incisor, no canine, three premolars, and three molars. The lower second premolar is single-rooted, and the third premolar is premolariform. The fourth lower premolar has a distinct metaconid, no paraconid, and a two-cusped talonid with a more fully-developed basin than in the closely related Arctodontomys. Lower molars each have a small yet distinct paraconid, a semi-compressed trigonid, a developed mesoconid, and a small, twinned hypoconulid. [9]
The upper canine of Microsyops is double-rooted. The upper fourth premolar has a distinct metacone and a weak parastyle. Upper molars exhibit clear conules, in particular a distinct metaconule, unlike the condition in the closely related Craseops. [9] Additionally, Microsyops upper molars lack a postprotocingulum, in contrast to the condition found in most early Paleogene primates. [2]
The basicranium of Microsyops has been described in detail in order to determine its affinities with respect to other mammals. Based on basicranial features, the internal carotid artery which supplied blood to the brain of Microsyops was primitive with respect to both extinct and extant euarchontans. These features include a transpromontorial groove indicating an unreduced internal carotid artery and grooves marking the course for both stapedial and promontorial branches of the internal carotid artery (Silcox et al. 2020). Another characteristic that suggests Microsyops was primitive is the presence of unexpanded caudal and rostral tympanic petrosal processes. [11] Unlike other plesiadapiforms, Microsyops lacked a bony auditory bulla (Gunnel 1989, Silcox et al. 2020). Additionally, Microsyops lacks the specialized cranial morphology considered characteristic of crown scandentians and dermopterans. [11]
The most characteristic aspect of the cranial morphology of Microsyops is the presence of a postorbital process. [2] [11] This trait is unlike the condition found in early Paleogene primates, which possess a full postorbital bar. However, Microsyops also differs from other plesiadapiforms, which lack either a postorbital bar or process. [11] The postorbital process of Microsyops has been described as being superficially similar to that of dermopterans. [2]
Due to limited available material, very little is known about the postcranial morphology of Microsyops, and Microsyopidae in general. [2]
Microsyops was first described by Joseph Leidy in 1872. He compared lower jaw fragments, found by Dr. J. V. Carter in the Bridger Basin of southwestern Wyoming, to the condylarth Hyopsodus gracilis, named by Professor O. C. Marsh of Yale University. At the time he believed the fragments to represent the same animal and proposed the new binomial combination of Microsyops gracilis. Leidy later compared his M. gracilis to Marsh's Limnotherium elegans, which was originally described as a diminutive mammal and later as a primate. [12] He concluded they were the same but with L. elegans as a species of the genus Microsyops, and that his original Microsyops gracilis should be properly named Microsyops elegans. [9]
Microsyopinae and Uintasoricinae are subfamilies within the plesiadapiform family Microsyopidae. Microsyops is a genus of the subfamily Microsyopinae which also includes the genera Arctodontomys, Megadelphus and Craseops. This subfamily includes the larger microsyopids. The subfamily Uintasoricinae includes the diminutive taxa Niptomomys, Uintasorex, and Choctawius. Microsyopidae is one of the longest-lived groups of plesiadapiforms, lasting 20 million years in North America from the late Paleocene to late Eocene (Silcox et al. 2021). Two families of plesiadapiforms, Microsyopidae and Paromomyidae, have representative taxa from the Uintan Land-Mammal Age (middle Eocene) while the Plesiadapidae and Carpolestidae disappeared at the end of the Paleocene. [13]
Recognized species of Microsyops includes M. elegans, M. annectens, M. scottianus, M. augustidens, M. kratos, M. latidens, M. cardiorestes, M. vicarius, and M. knightensis, with M. elegans being the type species. [13]
Some authors argue that microsyopids are plesiadapiforms while others suggest a dermopteran grouping. [14] However, the overall relationship between plesiadiforms and other living and fossil members of Euarchontoglires has been disputed. In a cladistic analysis including postcranial, cranial, and dental characteristics by Bloch et al. (2007), [15] microsyopids were found to be plesiadapiforms more distantly related to euprimates than plesiadapoids or paromomyoids, and without any special relationship to dermopterans. [14] However, while analyses support a euarchontan grouping, specific relationships of microsyopids to other plesiadapiforms, euprimates, scandentia, and dermoptera remain unresolved. [11] Microsyopids are generally thought to be euarchontans, and some researchers consider them to be stem primates. [2]
Microsyopidae lived from the late Paleocene to the middle Eocene in North America. [14] Conditions of the Eocene supported extensive subtropical woodland and rainforest environments which facilitate arboreal lifestyles. This time is also characterized by the Paleocene-Eocene Thermal Maximum, displaying the highest temperatures of the Cenozoic period. From this peak, steady temperature declines are displayed throughout the middle to late Eocene. [16]
A well-preserved skull of Microsyops annectens from Carter Mountain in northwestern Wyoming has been used to generate a virtual endocast via micro-CT. [14] Cranial capacity has been estimated as 5.9 cm3, yielding an encephalization quotient (EQ) of 0.26-0.52 depending on different body mass estimates and the choice of equation used to estimate EQ. Microsyops has larger EQ than Plesiadapis cookei , and falls in the lower range of estimates for early Paleogene primates. However, basicranial anatomy is remarkably primitive, because the auditory bulla was not ossified and there are only grooves, rather than bony tubes, for the intrabullar parts of the internal carotid artery and its dependencies. [11] The basicranial anatomy of Microsyops appears to be little changed from that of primitive placental mammals.
Colugos are arboreal gliding mammals that are native to Southeast Asia. Their closest evolutionary relatives are primates. There are just two living species of colugos: the Sunda flying lemur and the Philippine flying lemur. These two species make up the entire family Cynocephalidae and order Dermoptera.
Purgatorius is a genus of seven extinct eutherian species typically believed to be the earliest example of a primate or a proto-primate, a primatomorph precursor to the Plesiadapiformes, dating to as old as 66 million years ago. The first remains were reported in 1965, from what is now eastern Montana's Tullock Formation, specifically at Purgatory Hill in deposits believed to be about 63 million years old, and at Harbicht Hill in the lower Paleocene section of the Hell Creek Formation. Both locations are in McCone County, Montana.
Plesiadapis is one of the oldest known primate-like mammal genera which existed about 58–55 million years ago in North America and Europe. Plesiadapis means "near-Adapis", which is a reference to the adapiform primate of the Eocene period, Adapis. Plesiadapis tricuspidens, the type specimen, is named after the three cusps present on its upper incisors.
Plesiadapiformes is an extinct basal pan-primates group, as sister to the rest of the pan-primates. The pan-primates together with the Dermoptera form the Primatomorpha. Purgatorius may not be a primate as an extinct sister to the rest of the Dermoptera or a separate, more basal stem pan-primate branch. Even with Purgatorius removed, the crown primates may even have emerged in this group.
Shoshonius is an extinct genus of omomyid primate that lived during the Eocene. Specimens identified as Shoshonius have been found exclusively in central Wyoming and the genus currently includes two species, Shoshonius cooperi, described by Granger in 1910, and Shoshonius bowni, described by Honey in 1990.
Altanius is a genus of extinct primates found in the early Eocene of Mongolia. Though its phylogenetic relationship is questionable, many have placed it as either a primitive omomyid or as a member of the sister group to both adapoids and omomyids. The genus is represented by one species, Altanius orlovi, estimated to weigh about 10–30 g (0.35–1.1 oz) from relatively well-known and complete dental and facial characteristics.
Plesiadapidae is a family of plesiadapiform mammals related to primates known from the Paleocene and Eocene of North America, Europe, and Asia. Plesiadapids were abundant in the late Paleocene, and their fossils are often used to establish the ages of fossil faunas.
Carodnia is an extinct genus of South American ungulate known from the Early Eocene of Brazil, Argentina, and Peru. Carodnia is placed in the order Xenungulata together with Etayoa and Notoetayoa.
Mixodectidae is an extinct family of insectivorous placental mammals in the order Dermoptera. The mixodectids originated in the late Cretaceous and survived into the Paleocene in Europe and North America.
Plesiopithecus is an extinct genus of early strepsirrhine primate from the late Eocene.
The Willwood Formation is a sedimentary sequence deposited during the late Paleocene to early Eocene, or Clarkforkian, Wasatchian and Bridgerian in the NALMA classification.
Ocepeia is an extinct genus of afrotherian mammal that lived in present-day Morocco during the middle Paleocene epoch, approximately 60 million years ago. First named and described in 2001, the type species is O. daouiensis from the Selandian stage of Morocco's Ouled Abdoun Basin. A second, larger species, O. grandis, is known from the Thanetian, a slightly younger stage in the same area. In life, the two species are estimated to have weighed about 3.5 kg (7.7 lb) and 10 kg (22 lb), respectively, and are believed to have been specialized leaf-eaters. The fossil skulls of Ocepeia are the oldest known afrotherian skulls, and the best-known of any Paleocene mammal in Africa.
Micromomyidae (Micromomids) is a family of extinct plesiadapiform mammals that include some of the earliest known primates. The family includes five genera that lived from the Paleocene epoch into the early Eocene epoch.
Ignacius is a genus of extinct mammal from the early Cenozoic era. This genus is present in the fossil record from around 62-33 Ma. The earliest known specimens of Ignacius come from the Torrejonian of the Fort Union Formation, Wyoming and the most recent known specimens from Ellesmere Island in northern Canada. Ignacius is one of ten genera within the family Paromomyidae, the longest living family of any plesiadapiforms, persisting for around 30 Ma during the Paleocene and Eocene epochs. The analyses of postcranial fossils by paleontologists suggest that members of the family Paromomyidae, including the genus Ignacius, most likely possessed adaptations for arboreality.
Azygonyx was a small tillodont mammal, likely the size of a cat to raccoon, that lived in North America during the Paleocene and Eocene in the early part of the Cenozoic Era. The only fossils that have been recovered are from the Willwood and Fort Union Formations in the Bighorn Basin of Wyoming, United States, and date to the Clarkforkian to Wasatchian, about 56 to 50 million years ago. Fifty-six collections that have been recovered thus far include the remains of Azygonyx. Azygonyx survived the Paleocene Eocene Thermal Maximum along with other mammals like Phenacodus and Ectocion, both of which were ground-dwelling mammals. Azygonyx probably was a generalist terrestrial mammal that may have roamed around the ground, but was also capable of climbing trees.
Apatemys is a member of the family Apatemyidae, an extinct group of small and insectivorous placental mammals that lived in the Paleogene of North America, India, and Europe. While the number of genera and species is less agreed upon, it has been determined that two apatemyid genera, Apatemys and Sinclairella, existed sequentially during the Eocene in North America. The genus Apatemys, living as far back as 50.3 million years ago (mya), existed through part of the Wasatchian and persisted through the Duchesnean, and Sinclairella followed, existing from the Duchesnean through the Arikareean. Examinations of specimens belonging to the genus Apatemys suggest adaptations characteristic of arboreal mammals.
Carpodaptes was a genus that encompassed small, insectivorous animals that roamed the Earth during the Late Paleocene. Specifically, Carpodaptes can be found between the Tiffanian and Clarkforkian periods of North America. Although little evidence, this genus may have made it through to the early Eocene. They are known primarily from collections of jaw and teeth fragments in North America, mainly in southwestern Canada and northwestern America. Carpodaptes are estimated to have weighed approximately 53-96 grams which made them a little bigger than a mouse. However small, Carpodaptes was a placental mammal within the order Plesiadapiformes that appeared to have a high fiber diet. This insect-eating mammal may have been one of the first to evolve fingernails in place of claws. This may have helped them pick insects, nuts, and seeds more easily off the ground than with paws or claws. Carpodaptes was thought to only exist in North America but recent discoveries of dentition fragments have been found in China.
Navajovius is an extinct genus of plesiadapiforms that lived during the Paleocene epoch. Plesiadapiforms were small, arboreal mammals that are theorized to be either closely related to primates or dermopterans. Navajovius has only been documented from localities within North America. This genus was officially named in 1921 by Walter Granger and William Matthew and the type specimen is housed at the American Museum of Natural History.
Saxonella is a genus of extinct primate from the Paleocene Epoch, 66–56 Ma. The genus is present in the fossil record from around ~62–57 Ma. Saxonella has been found in fissure fillings in Walbeck, Germany as well as in the Paskapoo Formation in Alberta, Canada. Saxonella is one of five families within the superfamily Plesiadapoidae, which appears in the fossil record from the mid Paleocene to the early Eocene. Analyses of molars by paleontologists suggest that Saxonella most likely had a folivorous diet.
Torrejonia is a genus of extinct plesiadapiform that belongs to the family Palaechthonidae. There are currently two species known, T. wilsoni and T. sirokyi. This genus is present in the fossil record from around 62–58 Ma. Species belonging to this genus are suggested to be plesiadapiforms based on adaptations observed in the skeletal morphology consistent with arboreal locomotor behavior. Following the mass extinction event at the Cretaceous–Paleogene boundary (K-Pg), a large diversity of plesiadapiform families were documented beginning at the Torrejonian NALMA. Research has shown that T. wilsoni is one of the largest palaechthonids and is reconstructed as being more frugivorous than other palaechthonids.